Alternative Concepts of Reproductive Effort, Costs of Reproduction, and Selection in Life-History Evolution

An outline for an organismic theory of reproductive tacticsis presented to develop the demographic theory of optimal reproductivetactics into a more realistic theory of life-history evolution.Reproductive effort—denned as the proportion of resourcesinvested in reproduction—and the costs in somatic investmentdo not automatically result in survival costs. Both the conditionswhere survival costs are produced and the conditions where reproductioncan take place without survival costs are specified. Compensationand threshold hypotheses are put forward to allow weaker correlationsbetween reproduction and survival than the trade-off hypothesis,which assumes direct impacts by reproductive effort on survival.Furthermore, reproductive tactics are unlikely to be mouldedby the demographic forces of selection only. An empirical exampleis shown where residual reproductive value played no significantrole in the evolution of reproductive tactics. Selection probablyoperates not on separate life-history traits but on whole organismsthrough their entire life-history. The structural and physiologicalintercouplings between separate traits can result in phenotypicopportunity sets where selection can mould life-history traitsonly within the constraints of the opportunity sets. Optimizationtheory has provided an efficient technique for modelling andmaking predictions. However, organismic selection does not necessarilyoptimize adaptive strategies but eliminates unfit strategies.Life-history theory, and evolutionary theory in general, canbe developed along alternative logical lines when differenthypotheses are generated on how selection operates.     

The Hydraulic Principle

SYNOPSIS. AS theory rules method, the methodological proceduresapplied to morphological explanation have to be derived fromthe law-like properties of the objects under investigation.The explanation of organismic constructions has to be basedon the hydraulic principle which describes organisms as systemscomposed of fluid contained within flexiblemembranes. This insightestablishes a supra-molecular causal principle which, in itsgenerality for morphological explanation, parallels the biochemicalprinciples of molecular biology on the molecular level. Everyform and architectural arrangement has to be conceived as theresult of the form-enforcing influence of mechanical elementsthat operatewithin an integrated mechanically coherent system.An adequate explanation of morphological configuration has toelaborate the organization of the constructional whole and explainits properties as the result of a gradual transformation processthat is constrained by internal mechanical principles. The theoriesdeveloped by such a procedure are open to criticism and canbe tested and corroborated by reference to experiments conductedby nature.     

An organismic critique of molecular darwinism

The molecular darwinian approach to the emergence of life treats the competition between RNA sequences for nucleotide resources as the primordial selective process in prebiotic evolution, which prescribes possible pathways for the subsequent elaboration of organizational relationships. Since success in this competition is determined by the "phenotypic" properties of RNA strands in the absence of organizational context, the genesis of biotic organization is dependent upon the establishment of co-operative, hypercyclic interactions between competing RNA sequences. The thesis of this paper is that hypercycle theory is based on unwarranted assumptions about the conditions of prebiotic evolution, and that the implications of these assumptions run counter to both empirical evidence and to the rational by which natural selection operates in evolution generally. An organismic alternative to hypercycle theory is suggested, based on the catalytic microsphere and the thermodynamics of selection.     

A comparative analysis of the Darwin-Wallace papers and the development of the concept of natural selection

The classical theory of descent with modification by means of natural selection had no mother, but did have two English fathers, Charles Darwin (1809–1882) and Alfred Russel Wallace (1823–1913). In 1858,the Linnean Society of London published two contributions of these naturalists and acknowledged both authors as the proponents of a novel hypothesis on the driving force of organismic evolution. In the present report the most important sections of the Darwin-Wallace papers are summarized. This close reading of both publications reveals six striking differences in emphasis: Darwin and Wallace did not propose identical ideas. The species definitions of both authors are described and the further development of the concept of natural selection in wild populations is reviewed. It is shown that the contributions of A.R. Wallace, who died 90 years ago, are more significant than usually acknowledged. I conclude that natural selection's lesser known co-discoverer should be regarded as one of the most important pioneers of evolutionary biology, whose original contributions are underestimated by most contemporary scientists.     

The logical structure of irreversible systems transformations: A theorem concerning Dollo's law and chaotic movement

The argument of historical irrevocability as an explanation for the irreversibility of organismic evolution is reconsidered. It is examined under which conditions a chain of transformations never can lead to a state equivalent to a previous one. It is shown to be sufficient to take into consideration that the environment of an organ also consists of the other organs in an organism. Under this prerequisite it is impossible to reach an ancestral state once more. This type of irreversibility depends solely on the mutual interdependency of characters within one organism. No reference to statistical arguments is necessary. The relationship between chaotic dynamic systems and the present theory is discussed.     

Macroevolution: macrogenesis and typogenesis

One can distinguish two levels (and stages) of macroevolutionary processes: the lower (macrogenesis) and higher (typogenesis) ones. The macrogenesis represents macroevolutionary alterations of separate structures; the typogenesis is the forming of general Bauplan (type of organization) of a new macrotaxon on a base of initial macrogenesis. Discrete (or quantum) character of many macroevolutionary transformations is caused by various mechanisms, which are based on properties of integrated organismic systems and are characterized by threshold effect of their action. Initial macrogenesis can be resulted from the morphofunctional preadaptations; the pattern (or regime) transformations of morphofunctional organismic systems; emerging of dichotomy of morphogenetic programs and their following switching; the ontogenetic heterochronies (in particular, paedomorphosis); the allometric structural changes (and possibly some other mechanisms). The initial macrogenesis forms a base for qualitatively new adaptation and essentially influences on other systems in whole organism. That changes the selection vectors significantly. All these alterations trigger the typogenesis. The latter represents a complex of organismic systems transformations, integrated by selection and interconnections of various systems in whole organism. The important role in typogenesis belongs to the key alterations of some limiting organismic system that trigger and direct evolutionary changes of depended organismic systems. In course of typogenesis evolution, new macrotaxon occupies new adaptive zone.     

Objectivistic views in biology: an obstacle to our understanding of self-organisation processes in aquatic ecosystems

• 1 Ecologists are frequently inspired by the mechanistic view of classical physics in which motion is reduced to the effect of external forces on defined states of observable objects. Accordingly, dynamic events in ecosystems are resolved into environmental influences acting upon given states of an organismic system. In this conceptional scheme, both the environmental influences and the organismic system are assumed to be describable by objectively determinable parameters.
• 2 The present article directs criticism at this objectivistic approach. Accordingly, it is shown that an objectivistic view of living systems does not account for the complexity of organismic interactions which are continuously modified in a directional manner to achieve certain end states. This is exemplified in the physiological adaptation of micro‐organisms to their abiotic and biotic environment. Here, a population of single cells tends towards a ‘multicellular organism’ in which energy utilisation is optimised. During an investigation of this physiological adaptation process the organisms also adapt to the imposed experimental conditions, rendering futile any analysis in mechanistic terms.
• 3 Owing to this property of physiological adaptation a distinct research strategy is called for to establish the nature and ecological significance of this directionality. This strategy must be based on the observation that the sensitivity of living systems to a given environmental stimulus depends on the organismic prehistory with respect to previous exposures to stimuli. Thus, from an analysis of the adaptive response of a natural population to a defined challenge, information about prior environmental conditions may be derived that could not be obtained by other means.
• 4 Examples for the application of this research strategy to environmental problems are given.

     

Discussion: Leo Buss's The Evolution of Individuality

In his book The Evolution of Individuality, Leo Buss attacks a central dogma of the neo-Darwinian (or synthetic) theory of evolution, the idea that the individual is the sole unit of selection, by arguing that individuals themselves emerged as the result of selective forces that regulated the replication of cell lineages for the benefit of the whole organism. Buss also argues that metazoan developmental patterns and life cycles are the products of selection operating on different units of selection, and that there have been transitions between different units of selection during the history of life. Despite the revolutionary character of this book, The Evolution of Individuality in many ways reflects the adaptationist thinking often associated with the synthetic theory. Buss' framework could be improved by giving further consideration to chance factors in the evolution of development, and examining the details of the evolution of ontogeny in more depth.I am grateful to an anonymous reviewer from Biology and Philosophy for helpful comments and criticism.     

Maynard Smith,optimization, and evolution

Maynard Smith’s defenses of adaptationism and of the value of optimization theory in evolutionary biology are both criticized. His defense does not adequately respond to the criticism of adaptationism by Gould and Lewontin. It is also argued here that natural selection cannot be interpreted as an optimization process if the objective function to be optimized is either (i) interpretable as a fitness, or (ii) correlated with the mean population fitness. This result holds even if fitnesses are frequency-independent; the problem is further exacerbated in the frequency-dependent context modeled by evolutionary game theory. However, Eshel and Feldman’s new results on “long-term” evolution may provide some hope for the continuing relevance of the game-theoretic framework. These arguments also demonstrate the irrelevance of attempts by Intelligent Design creationists to use computational limits on optimization algorithms as evidence against evolutionary theory. It is pointed out that adaptation, natural selection, and optimization are not equivalent processes in the context of biological evolution. It is a pleasure to dedicate this paper to the memory of John Maynard Smith. Thanks are due to James Justus and Samir Okasha for comments on an earlier draft.     

Host defenses to transposable elements and the evolution of genomic imprinting

Genomic imprinting is the differential expression of maternally and paternally inherited alleles of specific genes. Several organismic level hypotheses have been offered to explain the evolution of genomic imprinting. We argue that evolutionary explanations of the origin of imprinting that focus exclusively on the organismic level are incomplete. We propose that the complex molecular mechanisms that underlie genomic imprinting originally evolved as an adaptive response to the mutagenic potential of transposable elements (TEs). We also present a model of how these mechanisms may have been co-opted by natural selection to evolve molecular features characteristic of genomic imprinting.     

What is this stuff called fitness?

This paper considers a variety of attempts to define fitness in such a way as to defend the theory of evolution by natural selection from the criticism that it is a circular argument. Each of the definitions is shown to be inconsistent with the others. The paper argues that the environment in which an animal evolves can be defined only with respect to the properties of the phenotype of the animal and that it is therefore not illuminating to try to explain the phenotypic properties of the animal in terms of adaptation to an environment that is defined by those very properties. Furthermore, since there is no way that the environment can be defined independently of the presence of the animal there is no way that the quality of an animal can be assessed; and there can be no objective criteria by whichany form of selection can be carried out, therefore there can be no criteria by whichnatural selection can be carried out. It is proposed that fitness is nothing more than the production of offspring, that this is a phenotypic property like all the others, and if it is heritable then the offspring of the parents that produce the most offspring will themselves produce the most offspring, and that in principle it is impossible to account for this in terms of the other phenotypic properties of the fittest animals except by circular argument. Differential rates of reproduction are the causes of evolution and the phenotypic causes are strictly inexplicable.     

Organismic sets: Outline of a general theory of biological and social organisms

The discussion as to whether societies are organisms andvice versa has been going on for a long time. The question is meaningless unless a clear definition of the term “organism” is made. Once such a definition is made, the question may be answered by studying whether there exists any relational isomorphism between what the biologist calls an organism and what the sociologist calls society. Such a study should also include animal societies studied by ecologists. Both human and animal societies are sets of individuals together with certain other objects which are the products of their activities. A multicellular organism is a set of cells together with some products of their activities. A cell itself may be regarded as a set of genes together with the products of their activities because every component of the cell is either directly or indirectly the result of the activities of the genes. Thus it is natural to define both biological and social organisms as special kinds of sets. A number of definitions are given in this paper which define what we call here organismic sets. Postulates are introduced which characterize such sets, and a number of conclusions are drawn. It is shown that an organismic set, as defined here, does represent some basic relational aspects of both biological organisms and societies. In particular a clarification and a sharpening of the Postulate of Relational Forces given previously (Bull. Math. Biophysics,28, 283–308, 1966) is presented. It is shown that from the basic definitions and postulates of the theory of organismic sets, it folows that only such elements of those sets will aggregate spontaneously, which are not completely “specialized” in the performance of only one activity. It is further shown that such “non-specialized” elements undergo a process of specialization, and as a result of it their spontaneous aggregation into organismic sets becomes impossible. This throws light on the problem of the origin of life on Earth and the present absence of the appearance of life by spontaneous generation. Some applications to problems of ontogenesis and philogenesis are made. Finally the relation between physics, biology, and sociology is discussed in the light of the theory of organismic sets.     

Why don't zebras have machine guns? Adaptation, selection, and constraints in evolutionary theory

In an influential paper, Stephen Jay Gould and Richard Lewontin (1979) contrasted selection-driven adaptation with phylogenetic, architectural, and developmental constraints as distinct causes of phenotypic evolution. In subsequent publications Gould (e.g., 1997a,b, 2002) has elaborated this distinction into one between a narrow "Darwinian Fundamentalist" emphasis on "external functionalist" processes, and a more inclusive "pluralist" emphasis on "internal structuralist" principles. Although theoretical integration of functionalist and structuralist explanations is the ultimate aim, natural selection and internal constraints are treated as distinct causes of evolutionary change. This distinction is now routinely taken for granted in the literature in evolutionary biology. I argue that this distinction is problematic because the effects attributed to non-selective constraints are more parsimoniously explained as the ordinary effects of selection itself. Although it may still be a useful shorthand to speak of phylogenetic, architectural, and developmental constraints on phenotypic evolution, it is important to understand that such "constraints" do not constitute an alternative set of causes of evolutionary change. The result of this analysis is a clearer understanding of the relationship between adaptation, selection and constraints as explanatory concepts in evolutionary theory.     

Contribution to the theory of organismic sets,III

In line with previous studies on organismic sets, the division of all organismic sets intogeneral autotrophic and heterotrophic is introduced. The first produce their food themselves from some external source of energy, which in general may be an energy of any kind. The others use other organismic sets as the source of their food and energy. On earth we know only one kind of generalgeneral autotrophic organismic sets, namely, the autotrophic plants which use solar radiation as their source of energy and for production of their own food. It is shown why autotrophic animals do not exist on earth except as microorganisms like, e.g.,Euglena. A rigorous proof of the previously derived theorem that in an organismic set of ordern>1 no element can be completely specialized is given. It requires the introduction of new postulates. Finally, in considering the organic world as a whole, the notion of organismic sets ofmixed order is introduced.     

Zum konzept der »inneren« selektion: Stellungnahme zu einer evolutionstheoretischen kontroverse

Recently the concept of natural selection in Darwin’s sense has been criticized by some authors. It has been argued that this concept does not explain certain phenomena of evolutionary change, especially in the reach of macroevolution. Some biologists, therefore, demanded for evolution a new model of selection which focuses internal factors in phytogeny. — This paper is a brief discussion of some aspects of “internal” selection and its meaning in contemporary evolutionary biology. The argument of the paper is that evolution can only be explained by a theory taking cognizance of interactions between external and internal selective agencies. Such a theory would be a systems theory of evolution.     

The Role of the Principle of Object-relatedness in the Theory of Activity

The present state of the theory of activity (by which we mean the concept developed by Leont'ev and his school) undoubtedly deserves serious discussion. Such a discussion must not be reduced, as often happens, to a critique of this theory by representatives of other schools and currents in psychology that are based on different theoretical positions (i.e., to a criticism from without), on the one hand, or to an unconditional acceptance of all of its postulates by proponents of this theory, on the other. In my view, the kind of critique of this theory that is most relevant and acutely necessary in terms of both its prospects of development and the extent to which its perceptions accord with other psychological currents is of another sort, namely, a critique from within, by which I mean analysis of its own laws, of the internal logic of development of the theory, and of its internal contradictions, which define its present status and the dynamics of its development. Such an analysis should enable us to stimulate more effectively the internal mechanism of development and self-development of the theory of activity and ensure a transition from a period of stagnation that has produced nothing of note to a progressive evolution of its content and its explanatory power.     

The uniqueness of biological self-organization: challenging the Darwinian paradigm

Here we discuss the challenge posed by self-organization to the Darwinian conception of evolution. As we point out, natural selection can only be the major creative agency in evolution if all or most of the adaptive complexity manifest in living organisms is built up over many generations by the cumulative selection of naturally occurring small, random mutations or variants, i.e., additive, incremental steps over an extended period of time. Biological self-organization—witnessed classically in the folding of a protein, or in the formation of the cell membrane—is a fundamentally different means of generating complexity. We agree that self-organizing systems may be fine-tuned by selection and that self-organization may be therefore considered a complementary mechanism to natural selection as a causal agency in the evolution of life. But we argue that if self-organization proves to be a common mechanism for the generation of adaptive order from the molecular to the organismic level, then this will greatly undermine the Darwinian claim that natural selection is the major creative agency in evolution. We also point out that although complex self-organizing systems are easy to create in the electronic realm of cellular automata, to date translating in silico simulations into real material structures that self-organize into complex forms from local interactions between their constituents has not proved easy. This suggests that self-organizing systems analogous to those utilized by biological systems are at least rare and may indeed represent, as pre-Darwinists believed, a unique ascending hierarchy of natural forms. Such a unique adaptive hierarchy would pose another major challenge to the current Darwinian view of evolution, as it would mean the basic forms of life are necessary features of the order of nature and that the major pathways of evolution are determined by physical law, or more specifically by the self-organizing properties of biomatter, rather than natural selection.     

Towards an organismic concept of land plants: The marginal blastozone and the development of the vegetation body of selected frondose gametophytes of liverworts and ferns

An organismic concept of land plants is outlined, which is based on a synthesis of plant morphology and plant anatomy. The entire plant, the living unity, is conceived as the organism being subdivided into cells, which cannot be interpreted as organisms themselves in the sense of elementary organisms. The evolution of land plant tissue systems is discussed in the introductive chapter.To test the proposed concept, some frondose plants were selected from liverworts (Pellia epiphylla, Metzgeria furcata, Pallavicinia lyallii) and comparable fern gametophytes (Dryopteris filix mas, Vittaria lineata, Stenochlaena tenuifolia) and studied with respect to their organization and the principles of development. They all have an archetypic, two-dimensional, open construction, which is described as the repens-type of plant construction. Primary form growth occurs in the marginal blastozone, which controls cell wall integration. One of the most significant processes of form generation is blastozone fractionation. The tissues leaving the blastozone differentiate during extension growth and maturation of the vegetation body. While the plant grows continuously in the blastozone, it decays steadily in the necrozone.The implications of the two-dimensional repens-type are discussed. It appears as a perfect plant construction, fit to start plant evolution on the land surface. Growing upwards into the atmosphere, the repens-type is obscured. But is reappears in all groups of higher land plants. This demonstrates the existence of evolutionary cycles in plants. It is argued that mutation and selection do not suffice to understand cyclical evolutionary patterns. The influence of organismic construction seems to predetermine evolution because of the limited options to change an appropriately functioning construction. Via construction analysis evolutionary options can be detected and thus, evolution becomes predictable to some extent. Instead of being object of mutation and selection, living organisms should be conceived as subjects in evolution (Weingarten 1993).Dedicated to my admired teacher Professor DrWilhelm Troll on the occasion of his 100^th birthday, 3rd November, 1997.